Seed collection and germination strategies for common wetland and coastal sage scrub species in Southern California.
Current estimates affirm that over 70% of coastal wetlands in the Southern California Bight have been lost since the 1800's, with estimates increasing to over 95% for highly urbanized areas, such as Los Angeles County (Stein et al. 2014). The magnitude of these losses and the continued degradation of coastal wetland systems, and adjacent upland and coastal sage scrub habitats, threatens the ecological integrity and sustainability of these habitat types and their watersheds. To address these issues, a number of restoration and mitigation projects aimed at restoring lost ecosystem services, increasing biodiversity, boosting resilience, and in the case of mitigation, creating new wetland habitat, are currently in the planning process in southern California (Noss 2000, Zedler 2000). The majority of wetland restoration or mitigation projects develop a site-specific framework of protocols and management strategies outlining a planting and re-vegetation strategy.
Planting strategies designed to establish self-sustaining plant communities identify both the species to be included in the restoration and the source of plant material (i.e. nursery stock or local seeds) (Zedler 2001). Restoration plant palettes should be designed to mimic reference or historic site diversity and be composed of an appropriately broad range of species (Zedler 2001, Johnston et al. 2012). Because of their unique location in the landscape as the connecting habitat between marine, terrestrial, and freshwater ecosystems, coastal wetland complexes naturally support a variety of salt marsh, brackish, and freshwater plant species (Lichvar et al. 2014). Species from each of these habitat types should be incorporated into an appropriate plant palette. Evidence also suggests that the wetland-upland ecotone should be considered an extension of wetland habitat for conservation and restoration purposes (James and Zedler 2000, Wasson and Woolfolk 2011). Thus, coastal sage scrub, dune, and transitional species commonly found in the wetland-upland ecotone should be considered in wetland restoration re-vegetation strategies.
Once a restoration plant palette has been developed, species-specific plant material (e.g. seeds and seedlings) acquisition and propagation methods must be determined. While plant material can be obtained from local nurseries, collection and propagation of native seed from local sites is considered the most cost-effective and ecologically-sound method of sourcing germplasm for restoration and mitigation projects (Zedler 2001, Broadhurst et al. 2008). Site-specific or nearest neighbor collections are preferred to distant collections and use of nursery stock, as locally-collected individuals are better adapted to community environmental conditions, maintain local genetic integrity, ensure persistence of local eco-types, prevent unintended gene flow, may improve the long-term sustainability of the site, and may enrich the diversity of the wetland plant community (Guerrant 1996, Montalvo et al. 1997, Bowler 2000, Zedler 2001, Mitsch and Gosselink 2010, Vander Mijnsbrugge et al. 2010). Non-local genotypes may be maladapted to local site conditions, leading to improper establishment, or negative impacts to plant and animal communities through competition or species hybridization (Bischoff et al. 2006, Vander Mijnsbrugge et al. 2010). The retention of local eco-types and genetic information is gaining importance in the field of restoration biology, reflected by the recent inclusion of required onsite and/or near neighbor collections by regulatory agencies overseeing restoration and mitigation work (Bowler 2000). It is also important to note that making collections in many nature preserves requires permits and/or express permission from the regulating agency.
A number of techniques exist to propagate plant material for wetland, coastal sage scrub, and dune species. Seeds are often the primary means of reintroducing native plant species to restoration sites in a number of habitat types (Montalvo et al. 2002, Merritt and Dixon 2011). Restoration sites may be seeded using a variety of techniques (e.g. broadcast seeding, drilling, imprinting, or hydroseeding) or collected, cultivated in a greenhouse, and transplanted to the site (Bowler 2000, Montalvo et al. 2002, Merritt and Dixon 2011). Simple seeding experiments generally are performed with limited success, especially at lower elevations or within tidal wetland habitats, as seeds often fail to germinate or float away with rising tides (Broome et al. 1988, Zedler 2001). Techniques like hydroseeding that involve mixing seed with water and either mulch, soil, or organic matter prior to application, tend to work well for many wetland and coastal sage scrub species [e.g. Salvia mellifera (black sage) and Eriogonum fasciculatum (California buckwheat)] and may enhance seedling establishment (Zedler 2001, Montalvo et al. 2002, Montalvo and Beyers 2010).
Transplanting greenhouse-grown seedlings is an effective re-vegetation strategy that may increase the potential establishment success when compared to direct seeding for some species. In one experiment, survivorship of 2-4 month old marsh seedling transplants was over 95% for all but one treatment, much higher than the success rate of direct seeding (Zedler 2001). Seedlings of a variety of halophytic marsh species including Suaeda esteroa, estuary seablite, and Salicornia bigelovii, dwarf pickleweed, and a variety of coastal sage scrub species like Atriplex canescens, four-wing salt bush, have been successfully grown in greenhouses and transplanted for restoration purposes (Zedler 2001, Francis 2009). While use of seeds and seedlings has been successful for many species [e.g. Achillea millefolium (common yarrow) and Astragalus tener var titi (coastal dunes milk vetch)], effective propagation techniques are species-specific and other species, like Batis maritime, saltwort, do not readily grow from seed and require use of alternate methods (Zedler 2001).
Other common approaches to generate plant stock include use of cuttings, root division, and direct transplantation of seedlings or mature plants to the site of interest (Zedler 2001, Baskin and Baskin 2014). Direct transplantation of coastal sage scrub seedlings [e.g. Artemisia californica (California sagebrush), Salvia mellifera, Encelia californica (California brittlebush), and Eriogonum fasciculatum] and mature plants salvaged from donor sites have been used with great success in mitigation efforts (Bowler et al. 1994, Bowler 2000). Similarly, use of transplants, sod, and small plugs of wetland soil, have been effective in introducing a number of wetland species, including Spartina foliosa, California cordgrass, to sites (Tmka 1998, Zedler 2001, Mitsch and Gosselink 2010). Use of cuttings is documented to work well for other species; cuttings of Salicornia pacifica, common pickleweed, for example, have been successfully propagated by Tree of Life Nursery in San Juan Capistrano, California. While each of these approaches has merit, the discussion in the remainder of this paper (and the accompanying appendices) focuses on the use of seeds and greenhouse-grown seedlings to target a data gap in peer-reviewed literature.
While general techniques for successfully establishing common wetland and coastal sage scrub species described in the preceding paragraphs are understood (Broome et al. 1988), the field of restoration biology is still developing and could benefit greatly from additional research. More specifically, the field could benefit from research regarding species-specific collection and propagation techniques because cultivation and planting strategies are often species-specific, highly variable, proprietary, or experimental. Information for many native species of interest does not exist, or is not publically available, forcing restoration managers and ecologists to rely on general information about the genus or costly and time-intensive exploratory studies (Dreesen and Harrington 1997). Publically available sources are scattered throughout a variety of peer-reviewed and non-peer-reviewed resources. With over a dozen wetlands in southern California considered candidates for large-scale wetland restoration projects, a compilation of literature summarizing re-vegetation strategies for the region is needed (SCCWRP 2001). This paper synthesizes basic seed characteristics, as well as collection and germination strategies for vegetation species common to estuarine wetland and adjacent upland habitat types, specifically coastal salt marsh and coastal sage scrub habitats in southern California.
Materials and Methods
Common seed collection, germination, and propagation techniques are described in the text of this paper. General species information (e g. scientific name, common name, and habitat type) is included in Appendix I. Detailed species-specific data and recommendations are included in Appendix II, which summarizes available information for 66 native plant species commonly used in southern California coastal restoration projects. Species-specific details were compiled using available literature. While the majority is derived from peer-reviewed publications, some non-peer reviewed literature was included to fill data gaps in published information. As many data gaps exist, and gray literature was used throughout the article text and the accompanying appendices, the authors have chosen not to distinguish gray literature with footnotes and this was approved by the editors. Instead, these sources are listed, with all peer-reviewed sources, in the Literature Cited section. In instances where duplicate information was identified, the source with the most extensive experimental results was cited. Field observations from the Ballona Wetland Ecological Reserve, Los Angeles, CA, were used to determine some seed collection windows. Appendix II is not intended to be comprehensive; instead, it focuses on common coastal wetland and upland species in southern California for which there was available literature. Priority was given to information specific to southern California coastal habitats, but species-specific information from other geographic areas was included as needed for completeness. Implementation of specific methods may vary slightly by site or project. A number of resources exist that provide general species profiles of the plants described in Appendices 1 and II. Three websites in particular, S&S Seeds (http://www.ssseeds.com), the Theodore Payne Foundation (http://theodorepayne.org), and Tree of Life Nursery (http://www.califomianativeplants.com), are recommended for supplemental information relating to life history and planting recommendations.
Equipment and supplies needed for seed collection, cleaning, and germination are highly variable based on the specific vegetation species. Recommended field, laboratory, and greenhouse equipment are listed in Table 1. In addition to the field equipment listed, available background information (e.g. reports, vegetation maps, taxonomic keys) should be brought into the field to aid correct taxonomic identification of species.
Seeds should be collected within seed zones, geographic zones in which genetic exchange naturally occurs. Practitioners are advised to use life history traits, landscape context, and available genetic studies to correctly determine seed zones (Krauss and He 2006). It is important to note that due to extensive urbanization and fragmentation in southern California, historic areas of seed exchange have been diminished. In addition to considering provenance of seeds, care should be taken to ensure that seed collections contain sufficient genetic diversity (Vander Mijnsbrugge et al. 2010) as diversity safeguards against disease, environmental fluctuations, and inbreeding depressions (Smith et al. 2007). To maximize the range of genetic diversity represented in the collection, seed should be collected from 10-50 individuals per population (Lippitt et al. 1994, Vander Mijnsbmgge et al. 2010). Local adaptations and site-specific variability should also be taken into consideration, but site-specific recommendations are outside the scope of this product. When collecting seeds, less intense and more frequent seed harvests are preferable to infrequent and intense harvests (Wall 2009). Negative impacts on the seed source population must be considered (Krauss and He 2006). A general safe harvesting recommendation is to take no more than 5% of seed from a given species and geographic area (Zedler 2001).
Once plant identity has been confirmed, carefully examine the seeds to assess maturity. Avoid collection of immature seed, as premature collection may result in low seed viability (Bonner and Karrfalt 2008, Baskin and Baskin 2014). In general, it is good practice to begin collecting seeds around the time that natural dispersal begins (Baskin and Baskin 2014). Seeds are considered ripe if seed capsules are dry and tan or brown in color, rather than yellow or green (Lippitt et al. 1994, Bonner and Karrfalt 2008, Baskin and Baskin 2014). Frequent visits to collection sites are suggested to repeatedly assess seed stage within the recommended collection time window. For species with insufficient published seed collection data or information, e.g. Artemisia douglasiana, detailed field notes are essential to pinpoint the ideal collection window and successfully collect seeds.
Once the seeds of target species are deemed ripe, the collection process can begin. Collection/ isolation of seed varies based on plant anatomy. Observe the plant and note if the species has berries or dry fruits, dehiscent or indehiscent seeds, and note if seeds are in seed heads or seed clusters as collection methods vary for each category (Table 2). Additionally, if a species is known to be dioecious [e.g. Croton californicus (California croton), Baccharis spp., Salix spp.], care should be taken to ensure that sufficient seed quantities are collected from both male and female plants (Clarke et al. 2007). Vouchering specimens from collected seeds is a good practice and should be considered during the planning phase.
Seed cleaning removes floral parts, seed coats, pods, fleshy fruit material, and other debris from seeds (Jorgensen and Stevens 2004). Machinery, including aspirators, hammermills, fanning mills, and blowers, exists to aid large-scale seed enterprises. Hammermills, fanning mills, and blowers help isolate seed and remove chaff and floral parts (Shaw 1975, Jorgensen and Stevens 2004). Although seed cleaning machinery is useful, cleaning for small-scale projects can be efficiently performed by hand (Bonner and Karrfalt 2008). To isolate seeds and remove excess chaff, remove seeds from branches and large floral parts. Then, rub remaining seeds and floral parts over a sieve. Once seeds are isolated from chaff, only retain seeds that look healthy and ripe (i.e. dark brown/tan in color, fully-formed). For some species, chaff does not present a huge problem, and it may be more efficient to seed with some chaff. Discard seeds that appear sickly or deformed. If the seed is contained in a capsule, gently crush the capsule by hand or with a rolling pin. Removal of woody capsules, as seen in Abronia spp., may also be aided with the use of generic nail clippers (RM. Drennan, personal communication).
For the greatest germination yield, storage time should be minimized, and use of newer seeds should be prioritized. While native seed longevity varies by genus and species, a number of seeds are known to be short-lived. For example, seeds of Lycium californicum, California box-thorn, are viable for up to one year at most. While seeds of other species [e.g. Atriplex spp., Astragalus spp., and Lupinus chamissonis (dune bush lupine)] will remain viable for much longer (i.e. 4-10 years), the germination rate of seeds in long-term storage will likely decline over time. In addition to reducing germination rate, long-term storage will often induce seed coat or embryo dormancy, and stored seeds may need to be treated prior to planting. For example, the hard seed coat of Astragalus tener var. titi seeds may require scarification, or mechanical scraping with sandpaper, a file, or a knife, to initiate germination if stored for an extended period of time (Baskin and Baskin 2014, USFWS n.d.)
The longevity of certain seeds can be increased if best management practices for storage are followed for the species and/or general seed storage procedures are applied. Most dry seeds should be stored at low temperatures, 10-15.6[degrees]C (50-60[degrees]F), and low humidity, less than 40% relative humidity (Jorgensen and Stevens 2004, Recon Native Plants Inc. 2015). Substandard storage in conditions with fluctuating temperatures or high humidity may result in significant seed loss (Merritt and Dixon 2011).
Successful propagation of southern California coastal plant species requires a thorough understanding of seed germination ecology. Seed germination is dependent upon a number of evolutionary and ecological factors which generally must be observed, and often replicated, in the laboratory or greenhouse to successfully grow propagules. These, often species-specific, factors include, but are not limited to: germination timing/seasonality, environmental conditions, such as temperature, soil texture, soil moisture, soil salinity, light availability, presence of smoke, and seed age, and dormancy state, both at the time of maturation and dispersal (Baskin and Baskin 2014).
Seeds are adapted to germinate under favorable environmental conditions (Deberry and Perry 2000). An understanding of natural germination timing is helpful in determining the environmental conditions that best promote germination of a particular species in the greenhouse or laboratory. This is particularly true, as in both the greenhouse and laboratory, environmental conditions can be manipulated to mimic natural seasonal variation. Temperature, moisture, and light are generally controlled for this purpose (see 'Temperature' and 'Light' sections below) (Noe and Zedler 2001).
Understanding germination timing under natural conditions will often indicate what range of temperatures best promote germination. Temperature influences germination directly through regulation of enzymatic reactions, or indirectly by controlling the synthesis of hormones that alter seed dormancy. While temperature is an important determinant in the regulation of both germination and dormancy, response to temperature in freshwater wetland species seems to be dependent on habitat, not phylogenetic relatedness. Temperature interplays with other environmental conditions to promote germination (Brandel 2006). Further, the germination rate of certain species is enhanced with simulated temperature fluctuations, rather than constant temperatures. While response to fluctuating temperatures depends both on specific species and habitat, a few generalities exist. Both small-seeded species and forbs tend to respond well to fluctuating temperatures while larger-seeded and graminoid species do not show as marked a preference for temperature fluctuations (Liu et al. 2013).
To grow seedlings, clean, viable seeds should be planted in mixtures of sand, top soil, and peat moss or vermiculite (Broome et al. 1988). To achieve the greatest germination rate, the exact composition of the mixture should be tailored to the individual plant species of interest. Life history and preferred habitat of the species should be considered when determining optimal soil conditions. For instance, Abronia maritima, which naturally occurs on sandy dunes, should be sown in soil consisting largely of sand, or other coarse grains.
Soil moisture must also be considered when sowing seeds (Noe and Zedler 2000, Noe and Zedler 2001). Most mature seeds must imbibe in the early stages of germination to activate enzymes (Deberry and Perry 2000). After seeds imbibe, sufficient, and relatively constant soil moisture is needed to ensure proper germination (Bonner and Karrfalt 2008). Most species in southern California salt marsh systems germinate well in moist soil at low salinity (Zedler 2001). Experiments suggest that Distichlis spicata grows best with a fluctuating inundation regime, where inundation was varied over time, but the soil surface was never completely dry (Elsey-Quirk et al. 2009). Germination of other high marsh plant species is highest with 41-51% soil moisture (Zedler 2001).
It is important to note that while seeds of wetland species are adapted to wet conditions with limited oxygen, coastal sage scrub and upland transition species are more sensitive to inundation. For these species, excessive exposure to water can be problematic, causing seeds to become waterlogged (Fenner 1992, Deberry and Perry 2000). Following germination, water regimes, that specify both the quantity and frequency of water application, both in the greenhouse and in natural environments, may influence growth rates and should be carefully considered.
Another major factor that influences germination is soil salinity (Noe and Zedler 2000, Noe and Zedler 2001). Certain halophytic species, like Salicornia bigelovii, germinate to higher percentages under somewhat saline conditions (0.05-0.09 M). In general, although halophytes are salt-tolerant, high percentages of halophyte seeds will germinate in distilled water. Results of salinity experiments suggest that seeds will often germinate to higher percentages in distilled water, as seeds tend to be sensitive to salt concentrations, and exposure to excessive salt can drastically decrease germination yields. Still, much variation exists in the germination of halophyte species in saline environments (Baskin and Baskin 2014).
Light is another environmental factor that affects germination. Exposure to light is often required for germination to occur. Exposure to light has been documented to improve germination rates for certain species [e.g. Eriogonum fasciculatum (California buckwheat), Baccharis salicifolia (mule fat)] (Zedler 2001, Bonner and Karrfalt 2008). Still, exposure is not always sufficient to ensure the successful occurrence of germination mechanisms. Duration of exposure to light (i.e. day length or photoperiod) also plays an important role in seedling emergence and growth of southern California natives (Sprague 1944, Noe and Zedler 2000, Greiner and Kohl 2014). For instance, long-day conditions (16 hours of light for every 8 hours of darkness) are necessary to successfully culture Oenothera species (Greiner and Kohl 2014). Photoperiod may also influence other processes, such as flowering. Melica imperfecta and Stipa lepida have been shown to flower 10-20 weeks faster with constant light (i.e. 24-hour photoperiod) when compared to an 8-hour photoperiod (Ashby and Flellmers 1959).
Southern California, like most regions with Mediterranean climates, is subject to frequent and intense wildfires, and certain species have adapted to be fire-tolerant (Keeley and Fotheringham 1998, Crosti et al. 2006). Germination of fire-tolerant species is generally enhanced by exposure to fire or smoke (Crosti et al. 2006, Baskin and Baskin 2014). Smoke-stimulated germination, via exposure to liquid or aerosol components of smoke, may be useful for many coastal sage scrub species. For instances, exposure of Salvia mellifera seeds to smoke or other components of fire, like charred wood or potassium nitrate (KN[O.sub.3]), may help stimulate germination (Montalvo and Beyers 2010).
In some instances, information regarding the necessary conditions or procedures to promote germination is not readily available for a particular species. In such situations, it is advisable to consult local experts that may have species-specific knowledge. Alternatively, simple tests or experiments manipulating a variety of the environmental factors discussed above may be performed.
If a seed lot requires germination studies, it is preferable that they are conducted shortly after seed collection, within 7-10 days, to ensure seeds are viable and have not entered seed dormancy. Germination trials can test outcomes of various pre-treatments and/or growing conditions. They are often also used to express the quality of a seed lot (Lippitt et al. 1994). The results of germination trials are typically reported as percentage germination or germination rates. Percentage gennination is the percentage of seeds that germinate under the specified set of conditions. Comparing germination rates of a variety of treatments allows easy determination of the most effective combination of germination conditions.
While germination rates are useful, the industry will often use other terms to describe the percentage of seed that will germinate under a given set of conditions. Pure Live Seed (PLS) is a common way to express viability. PLS is calculated by multiplying the percentage of pure seed by the percentage of total viable seed and dividing the product by one hundred (S&S n.d., Showers 2010). Other measures include specification by purity, bulk pounds, or PLS pounds (S&S).
Seeds for a number of wetland plants are known to be dormant. In these species, seed dormancy must be broken to promote growth and germination (Baskin and Baskin 2014). The process is generally moisture and temperature dependent, but varies both with species and type of dormancy. Three types of dormancy should be considered: physical (or seed coat) donnancy, internal dormancy, and morphophysiological dormancy. Seeds with physical dormancy have seed coats or other structures that are impermeable to water and/or oxygen (Lippitt et al. 1994, Baskin and Baskin 2014). This form of dormancy is generally broken by penetrating/ opening the seed coat or specialized structure that excludes water or oxygen. This can be achieved through scarification, cold and warm stratification, or exposure to dry heat, charate, fire, acid, and light. Internal dormancy, caused by a physiological mechanism that inhibits germination, is generally broken through use of warm and/or cold stratification. Morphophysiological dormancy is similar to physiological, but seeds with this type of dormancy also have an underdeveloped embryo. A variety of methods can be used to break morphophysiological dormancy, including: scarification, submersion in hot water [82-93[degrees]C (180-200[degrees]F)], treatment with dry heat, exposure to fire, acid, mulch treatment, cold stratification, warm stratification, and exposure to light (Emery 1988, McClure 1997, Baskin and Baskin 2014). Common dormancy breaking methods are detailed in Table 3).
Unfortunately, as indicated by the variety of conditions listed above, there is not one prevailing standardized method to break seed dormancy. Again, methods vary based on the life history of the species. Species-specific life histories, available at the growers' websites listed above, can be a good indicator of the required conditions for that species. For example, species that typically germinate in early spring after a cold and/or rainy winter, such as Platanus racemosa, western sycamore, often require cold, moist stratification mimicking natural wintering to break dormancy. Other species, such as Acmispon glaber, common deerweed, require heat treatment to break dormancy which also correspond with the life history of that species; A. glaber does particularly well after wildfire events. However, treating seeds to break dormancy is not enough to guarantee germination. Germination requirements must also be considered. Methods and information should be supplemented by experimentation when necessary.
Establishing functional ecosystems also requires consideration of subsurface components of the system. Many plants have symbiotic relationships with soil-inhabiting microorganisms, yielding root systems that are more effective at extracting water and nutrients from the rhizosphere (i.e. soil profiles influenced by root secretions and soil fauna). The fungus-root system is called mycorrhizae (Gerdemann 1968, Tree of Life Nursery n.d.). Research has shown that mycorrhizae can increase plant growth and are essential in successfully establishing vegetation during restoration and mitigation projects (Reeves et al. 1979, Allen and Allen 1980, Cooke and Lefor 1990). If planting areas are severely disturbed and lack a healthy rhizosphere, steps should be taken to ensure presence of mycorrhizae, or to increase the potential for natural development. As the presence of mycorrhizae is important in establishing many wetland and coastal sage scrub species, container plants are often inoculated prior to planting (Cooke and Lefor 1990, Bowler 2000). Seedlings can be inoculated with a spore suspension or via introduction of small amounts of collected soil from sites with a healthy rhizosphere to a sterile soil (van de Voorde et al. 2012). Starter-cultures are also available commercially.
Southern California has lost a significant portion of its coastal ecosystems due to urban development, agriculture, invasive species, and in the case of coastal estuarine wetlands, severely modified hydrology resulting from both channelization and deposition of fill sediments (Westman 1981). Loss of these ecosystems is concerning because they provide valuable ecosystem services including supporting important fisheries, filtering water, sequestering carbon, and providing habitat for a diversity of plant and animal life, including a number of threatened and endangered species. Wetlands are buffered by transition habitats, and many wetland-associated species also require adjacent upland habitat areas to breed, roost, or to have the highest likelihood of survival. Plant species in Southern California also display a high degree of endemism and the Southern California coast is considered a global biodiversity hotspot (SCCWRP 2001).
Although wetlands in southern California have attained protected status and efforts are being made to restore degraded habitats throughout the Southern California Bight, the increasing human populations along the California coast will continue to impact these coastal ecosystems (Callaway and Zedler 2004). To preserve the spectrum of ecosystem services coastal wetlands and their adjacent upland habitats provide, managers throughout the southern California region need to work collectively to conserve remaining high quality coastal wetland habitat and to restore lower quality, degraded habitats.
Clearly, there is a significant and ongoing regional need for restoration projects to recover lost habitats and preserve the unique communities. Increased reliance on ecological restoration of vegetation assemblages emphasizes the need for sound, scientifically-tested techniques to ensure the successful reestablishment of plant communities. While this document is not comprehensive, and there is still a practical need for land managers to compile detailed site information and evaluate site-specific experiments prior to implementing a restoration scheme, this literature review compiles available seed collection and germination information for the southern California region and provides an initial assessment of published methods for common wetland, dune, and coastal scrub plants. Many unknowns remain in restoration ecology theory, and understanding of the most effective restoration practices remains incomplete. Knowledge gaps regarding the collection and germination requirements of integral species [e.g. Hazardia squarrosa (saw-toothed goldenbush)] and other species with limited research available [e.g. Elymus triticoides (creeping wild rye)] precluded their inclusion in this review. Planners are encouraged to conduct regular site monitoring and employ adaptive management strategies. In this way, progress can be evaluated and unexpected outcomes and shortcomings can be corrected.
Still, there is a regional need for additional research regarding seed phenology and maturation of southern California species. Although a number of wetland, dune, and coastal sage scrub restorations are planned in southern California, information regarding seed collection and germination for many naturally occurring species is not readily available. Therefore, the field of plant community restoration could benefit greatly from additional research regarding seed phenology and maturation, both in the form of species-specific experimentation and literature and broader-scale, regional or ecosystem-based reviews. Filling in existing knowledge gaps and developing a better understanding of seed processes will help restoration ecologists collect high quality, viable seed, thereby increasing the potential success of the restored vegetation community by reducing seed/seedling mortality, restoration cost and human effort.
Perhaps more importantly, the region could benefit from the development of a coordinated network of restoration ecologists. Compilation of this literature review suggests that information regarding the restoration of wetland plant communities is abundant, but it is dispersed, produced by various sources, and often proprietary. Intentional withholding of information by nurseries or private environmental consulting firms inevitably leads to duplication of efforts by groups working in the southern California region and surely impacts both the overall quality of restored habitats and project efficiency. Engagement and cooperation of existing private industry groups and public sector regulators with a vested interest in restoring coastal wetland plant communities would be a major victory and a tangible step forward for the threatened coastal ecosystems in the region.
While establishing vegetation in restored wetlands is a vital component to the overall restoration scheme, it is just a small part of the overall restoration process. Restoring wetland ecosystems is complex; plans must incorporate vegetation, hydrology, substrate, and marine and terrestrial animals. To fulfill restoration aims, well-informed, inter-disciplinary approaches that incorporate ecologists, engineers, managers, lawyers, and practitioners from other technical fields are needed (Zedler 2000, Kiehl 2010). Inter-disciplinary approaches will best foster creativity and progress knowledge and understanding in the field of restoration.
The authors wish to thank the colleagues and interns who assisted with this work. Special thanks to Phillippa M. Drennan, Remy G. Landon, and Jessica Sharpe whose contributions were significant. Thanks as well to Recon Native Plants, Inc. This work was made possible with financial support from the California State Coastal Conservancy and the US Environmental Protection Agency, but does not necessarily reflect their views and policies.
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Appendix I. Species-specific habitat associations for wetland, coastal sage scrub, and upland transition species common in southern California. This table includes scientific and common names from Jepson eFlora (http://ucjeps.berkeley.edu/, accessed June 4, 2015). Habitat association information is derived from Jepson and further refined with information available from the Manual of California Vegetation (2nd edition), the S & S Seeds Plant Database (www.ssseeds.com/plant-database), and the species-specific literature cited in Appendix II (Baldwin et al. 2012, Sawyer et al. 2009, S&S Seeds, n.d.). Scientific name Common name Abronia maritima Red sand verbena Abronia umbel lata Pink sand verbena Achillea millefolium Common yarrow Acmispon glaber Deer vetch Ambrosia psilostachya Western ragweed Artemisia californica California sagebrush Artemisia douglasiana Mugwort Artemisia tridentata Big sagebrush Arthrocnemum subterminale Parish's pickleweed Astragalus pycnostachyus var. Ventura marsh milk vetch lanosissimus Astragalus tener var. titi Coastal dunes milk vetch Atriplex californica California orach A trip lex canescens Fourwing saltbush Atriplex lentiformis Big saltbush Atriplex prostrata Fat-hen Atriplex watsonii Watson's saltbush Baccharis pilularis Coyote brush Baccharis salicifolia subs, Mule fat salicifolia Baccharis sarothroides Broom baccharis Batis maritima Saltwort Cressa truxillensis Alkali weed Croton califomicus California croton Distichlis littoralis Shore grass Distichlis spicata Salt grass Encelia californica California brittlebush Eriogonum fasciculatum California buckwheat Frankenia salina Alkali heath Grindelia camporum Valley gum weed Hazardia squarrosa Saw-toothed goldenbush Heteromeles arbutifolia Toyon Hordeum brachyantherum Meadow barley Isocoma menziesii Coastal goldenbush Iva axillaris Poverty weed Jaumea camosa Fleshy jaumea Juncus acutus subs, leopoldii Southwestern spiny rush Juncus bufonius Toad rush Limonium californicum Western marsh-rosemary Lupinus chamissonis Dune bush lupine Lycium californicum California box-thorn Melica imperfecta Little California melica Mimulus aurantiacus Sticky monkey Oenothera elata Hookers evening primrose Peritoma arborea Bladderpod Phacelia ramosissima Branching phacelia Plantago erect a Foothill plantain Platanus racemosa Western sycamore Populus fremontii subs, fremontii Fremont cottonwood Potentilla anserina subs, pacifwa Pacific silverweed Pseudognaphalium californicum California cudweed Rosa californica California wild rose Salicomia bigelovii Dwarf pickleweed Salicomia pacifwa Common pickleweed Salix exigua subs, exigua Narrow-leaved willow Salix lasiolepis Arroyo willow Salvia apiana White sage Salvia mellifera Black sage Schoenoplectus acutus var. Common tule occidentalis Schoenoplectus califomicus Southern bulrush Spartina foliosa California Cord grass Stipa cemua Nodding needle grass Stipa lepida Foothill needle grass Suaeda esteroa Estuary seablite Suaeda nigra Bush seepweed Suaeda taxifolia Woolly seablite Triglochin maritima Common arrow-grass Vulpia microstachys var. Small fescue microstachys Low Mid High Salt Scientific name marsh marsh marsh pan Abronia maritima Abronia umbel lata Achillea millefolium Acmispon glaber Ambrosia psilostachya Artemisia californica Artemisia douglasiana Artemisia tridentata Arthrocnemum subterminale x x Astragalus pycnostachyus var. x x x lanosissimus Astragalus tener var. titi x x Atriplex californica x A trip lex canescens Atriplex lentiformis Atriplex prostrata x Atriplex watsonii x Baccharis pilularis Baccharis salicifolia subs, salicifolia Baccharis sarothroides Batis maritima x x x Cressa truxillensis Croton califomicus Distichlis littoralis x x Distichlis spicata x x Encelia californica Eriogonum fasciculatum Frankenia salina x x x Grindelia camporum Hazardia squarrosa Heteromeles arbutifolia Hordeum brachyantherum Isocoma menziesii Iva axillaris Jaumea camosa x x Juncus acutus subs, leopoldii Juncus bufonius x x Limonium californicum x x Lupinus chamissonis Lycium californicum Melica imperfecta Mimulus aurantiacus Oenothera elata Peritoma arborea Phacelia ramosissima Plantago erect a Platanus racemosa Populus fremontii subs, fremontii Potentilla anserina subs, pacifwa x x Pseudognaphalium californicum Rosa californica Salicomia bigelovii x x Salicomia pacifwa x x Salix exigua subs, exigua Salix lasiolepis Salvia apiana Salvia mellifera Schoenoplectus acutus var. occidentalis Schoenoplectus califomicus Spartina foliosa x Stipa cemua Stipa lepida Suaeda esteroa x x Suaeda nigra Suaeda taxifolia Triglochin maritima x x x Vulpia microstachys var. microstachys Low High Scientific name transition transition Grass Abronia maritima x Abronia umbel lata x Achillea millefolium x x Acmispon glaber x x Ambrosia psilostachya Artemisia californica Artemisia douglasiana x x Artemisia tridentata x Arthrocnemum subterminale Astragalus pycnostachyus var. x lanosissimus Astragalus tener var. titi x x Atriplex californica x x A trip lex canescens Atriplex lentiformis Atriplex prostrata x Atriplex watsonii x x Baccharis pilularis x Baccharis salicifolia subs, salicifolia Baccharis sarothroides Batis maritima x Cressa truxillensis Croton califomicus x x Distichlis littoralis Distichlis spicata x Encelia californica x Eriogonum fasciculatum Frankenia salina x x Grindelia camporum Hazardia squarrosa x Heteromeles arbutifolia x Hordeum brachyantherum x x Isocoma menziesii x x Iva axillaris Jaumea camosa Juncus acutus subs, leopoldii Juncus bufonius x Limonium californicum x Lupinus chamissonis Lycium californicum x x Melica imperfecta x Mimulus aurantiacus x Oenothera elata x Peritoma arborea Phacelia ramosissima x x Plantago erect a x Platanus racemosa Populus fremontii subs, fremontii Potentilla anserina subs, pacifwa x x Pseudognaphalium californicum Rosa californica x x Salicomia bigelovii Salicomia pacifwa Salix exigua subs, exigua Salix lasiolepis Salvia apiana Salvia mellifera Schoenoplectus acutus var. occidentalis Schoenoplectus califomicus Spartina foliosa Stipa cemua x Stipa lepida x Suaeda esteroa Suaeda nigra Suaeda taxifolia Triglochin maritima Vulpia microstachys var. x microstachys Fresh Salt Scientific name Scrub water tolerant Abronia maritima x Abronia umbel lata x Achillea millefolium x Acmispon glaber x x Ambrosia psilostachya Artemisia californica x Artemisia douglasiana x Artemisia tridentata x x Arthrocnemum subterminale x Astragalus pycnostachyus var. x lanosissimus Astragalus tener var. titi x x Atriplex californica x x A trip lex canescens x x Atriplex lentiformis x x Atriplex prostrata x Atriplex watsonii x Baccharis pilularis x Baccharis salicifolia subs, x salicifolia Baccharis sarothroides x x Batis maritima x Cressa truxillensis Croton califomicus x x Distichlis littoralis x Distichlis spicata x Encelia californica x Eriogonum fasciculatum x Frankenia salina x Grindelia camporum x x Hazardia squarrosa x x Heteromeles arbutifolia x x Hordeum brachyantherum x x Isocoma menziesii x x Iva axillaris Jaumea camosa x Juncus acutus subs, leopoldii Juncus bufonius x Limonium californicum x Lupinus chamissonis x Lycium californicum x Melica imperfecta x Mimulus aurantiacus x Oenothera elata x Peritoma arborea x x Phacelia ramosissima x x Plantago erect a x x Platanus racemosa x x Populus fremontii subs, fremontii x x Potentilla anserina subs, pacifwa x Pseudognaphalium californicum Rosa californica x x Salicomia bigelovii x Salicomia pacifwa x Salix exigua subs, exigua x Salix lasiolepis x Salvia apiana x x Salvia mellifera x x Schoenoplectus acutus var. x occidentalis Schoenoplectus califomicus x x Spartina foliosa x Stipa cemua x Stipa lepida x Suaeda esteroa x Suaeda nigra x Suaeda taxifolia Triglochin maritima x Vulpia microstachys var. x x microstachys Appendix II. Detailed, species-specific seed collection, seed germination, and seed storage information for wetland, coastal sage scrub, and upland transition species native to south-ern California. Information is sorted alphabetically by scientific name. Fruit/seed Scientific name Start End characteristics Abronia maritima May Aug Winged fruit 10-14 mm (Drennan 2008, Baldwin long. Fruit contains et al. 2012) single-seeded achenes. Abronia umbellate May Aug Winged fruit 6-13 mm (Drennan 2008, Baldwin long. Fruit contains et al. 2012, Center single-seeded achenes. for Plant Conservation 2015,) Achillea millefolium Aug Oct Oblong fruit, usually 2 (Baskin and Baskin mm in length, contains 2002a, Baldwin brown disk achenes. et al. 2012) Seeds mature in late summer-early fall. Acmispon glaber May Jul Narrow, bean-shaped, (Montalvo and Beyers curved seed pods 1-2 2010c) mm long. Indehiscent pods ripen in 4-6 weeks. Mature pods are dry and brown or olive green. Ambrosia psilostachya Oct Dec Brown bur 3-4.5 mm long (Pavek 1992, Baldwin contains tiny achenes. et al. 2012) Artemisia californica Oct Feb Fruit 0.8-1.5 mm long. (Hauser 2006, Young- Very small achenes, Mathews 2010, Baldwin generally mature in et al. 2012) early fall or winter; wind dispersed. Artemisia douglasiana <1 mm, glabrous fruit. Elkhom Slough National Small, ellipsoid, Estuarine Research hairless achenes Reserve 2001, without ribs or Shultz 2014) angles. Artemisia tridentate Sep early Fruit glandular or (Elkhom Slough winter hairy, 1-2 mm in National Estuarine length. Very small Research Reserve 2001, achenes, generally Baldwin et al. 2012, mature in early fall Tilley et al., n.d.) or winter. Arthrocnemum Oct Dec Stems have tiny flowers subterminale (Zedler that occur below the 2001, Clarke et al. tip of the stem and 2007, Baldwin et al. contain brown, 2012) vertical seeds with hard seed coat, 1-1.4 mm in length. Astragalus pycnostachyus Jul Fruit ovate and var. ianosissimus inflated; 6-11 mm long (McCue 2010, Baldwin and 3.5-6 mm wide. et al. 2012, U.S. Fish Seeds are smooth, and Wildlife Service, compressed with a n.d.) small notch at attachment site. 2 or more seeds/fruit. Astragalus tetter var. May Fruit 6-50 mm long and titi (Showers 2010, 1.7-3.5 mm wide. Seeds Baldwin et al. 2012) are smooth, compressed with a small notch at attachment site. 2 or more seeds/fruit. Atriplex calijarnica Sep Oct Mature fruit is an (Young 2001a) utricle with 1 seed. Seeds are black, shiny, hard, round, and flat; 2 mm at maturity. Atriplex canescens Oct Apr Cream-colored 4 winged (Springfield 1970, utricle, 5-23 mm wide. Baldwin et al. 2012) Seeds 1.5-2.5 mm long with brown, papery inner seed coat. Species has high percentage of empty seed. Smaller fruits tend to have higher percentages of filled seed. Atriplex lentiformis Sep Jan Produces large amounts (Young et al. 1980, of dark brown, 1.5 mm Baldwin et al. 2012) long seeds. Atriplex prostrated Sep Oct Two types of seeds: (Khan and Ungar 1984, brown, 1-2.5 mm long, Zedler 2001, Baldwin and black, 1-1.5 mm et al. 2012) long. Atriplex watsonii Jun Sep Seeds light brown, about (Zedler 2001, 1 mm long. Bryant 2004) Baccharis pilularis Aug Dec Single-sex white (Bonner and Karrfalt flowers. Glabrous, 2008, Montalvo et al. ribbed fruit 1-2 mm 2010b, Baldwin et al. long, pappus 5.5-9 mm 2012) long. Mature seeds are tiny, dark brown achenes with ring of long, unbranched pappus bristles. Baccharis salicifolia May Jul Glabrous, ribbed fruit subs. Salicifolia 0.8-1.3 mm long, (Bonner and Karrfalt pappus 3-6 mm long. 2008, Baldwin et al. Tiny achenes with a 2012) bristly pappus. Baccharis sarothroides Glabrous, ribbed fruit (Bonner and Karrfalt 2-2.6 mm long, pappus 2008, Baldwin et al. 2-3 mm long. Tiny 2012) achenes. Bads maritime Oct Nov Hard-walled lenticular (Zedler 2001, Marcone or oblong seeds. 1 mm 2003, Francis 2009, in length. Lonard et al. 2011) Cressa truxtilensis Jul Aug Fruits are small hairy (Elkhom Slough capsules, 1/8" long. National Estuarine Seeds pinkish in Research Reserve 2001, color, broadly Zedler 2001) egg-shaped. Croton californicus Jul Nov Flowers develop into (Young 2001b, Baldwin compact, greenish seed et al. 2012) pods. Mature seeds are smooth, round, and brown with tan spots. 3.5-5.5 mm in length. Distichlis lift oralis Jun Sep Spikelets, 8-13 mm in (Zedler 2001, Clarke length, generally et al. 2007, Baldwin concealed by leaves. et al. 2012) Seeds are quite small and remain in flowers until senescence. Distichlis spicata Sep Nov Spikelets 6-20 mm long. (Baskin and Baskin Seed likely dormant at 2002b, Elsey-Quirk time of dispersal. et al. 2009, Baldwin et al. 2012) Encelia californica Fruit 5-7 mm long, (Bonner and Karrfalt slightly longer with 2008, Baldwin et al. pappus. Seeds dark 2012) brown at maturity. Eriogonum fasciculatum May Aug Glabrous fruit 1.8-2.5 (Zedler2001, Montalvo mm in length. and Beyers 2010a, Baldwin et al. 2012) Fran ken ia salina Sep Oct Ellipsoid seed capsules (Young 2001k) (8 mm) contain 1 mm long, brownish black seeds. Ovular in shape with pointed tips. Grindelia camporum Jun Oct Small, long, and flat (Zafar and Shah 1994, achenes. Wind-borne, Bliss 2012) dandelion-like achenes with feathery tufts. Hazardia squarrosa Fruit: 5-8 mm, 5-angled, (Keil et al. 2013) glabrous; pappus 7-12 mm, white to red-brown in color. Oblong to lanceolate seeds. Heteromeles arbutifolia Oct Jan Large, smooth brown (Bonner and Karrfalt seeds. 2-3 seeds per 2008, Baldwin et al. pome. 2012, Gordon 2014, Recon Native Plants Inc. 2015) Hordeum brachyantherum Jun July Mature inflorescences (Elkhom Slough are light brown. National Estuarine Research Reserve 2001, Young 2001d) Isocoma menziesii Sep Nov Tan-colored achenes, (Zedler 2001, Wall and longer than wide, Macdonald 2009, wider on the plumose Montalvo and Beyers end, with lengthwise 2010b) striations. The top of the achene has a ring of white bristles. Seeds mature when the pappus becomes fluffy and achenes detach easily from the receptacle. Iva axillaris 1-2 seeds/head. 2 mm (Montalvo et al. long, turnip-shaped, 2010b) light, and buoyant. Jaumea carnosa Jul Oct Seeds are linear achenes (Young 2001e) with longitudinal stripes. Juncus acutus subs. Aug Nov Shiny brown capsules leopoldii contain multiple (Zedler2001, Baldwin irregularly shaped et al. 2012) seeds. Seeds can be narrowly winged. Juncus bufonius Mar May Ovoid or elliptic seeds. (Zedler 2001, Baldwin Seeds generally et al. 2012) 0.3-0.6 mm long. Limonium californicum Sep Nov 3 mm long narrow (Young 200If) ellipse, dark brown/ red at maturity. Lupinus chamissonis Apr Jun Hairy legume pods (Young 200 lg) 2.5-3.5 cm long. Mature seeds are dark brown and speckled and 3-4 mm in length. Lycium californicum Jan Feb 3-6 mm red berries. 2 (Zedler 2001, Baldwin oblong seeds per et al. 2012) berry. Melica imperfecta Apr Jun Mature inflorescences (Ashby and Hellmers are brown; mature 1959, Emery 1988, seeds are tan. Young 2001h, Baskin and Baskin 2002c) Mimulus aurantiacus Jun Aug Mature capsules are (Young 2001c, Baldwin brown and contain et al. 2012) tiny, black seeds less than 1 mm in length. Oenothera elata Seeds are irregularly (Greiner and Kohl shaped, stacked in 2014, B & T World small, brown, woody Seeds 2015, Dave's capsules with four Garden 2015a, chambers each with two Kleinman, n.d.) rows of small seeds. Peritoma arborea Capsules 3-6 cm long and (Lippitt et al. 1994, 1-2.5 cm wide. Mature Borders et al. 2008, fruits will often Baldwin et al. 2012) split at the seam, revealing seeds. Dark-colored seeds tend to be more viable that light-colored seeds. Phacelia ramosissima Capsules contain 8-12, (Baldwin et al. 2012) 1-2 mm long pitted seeds. Plantago erecta Apr 2-2.5 mm long. (Gulmon 1992, Montalvo et al. 2010a, Baldwin et al. 2012) Platanus racemosa Jun Spring Chestnut brown seed pods (Bonner and Karrfalt at maturity, many are 2008) empty. Achenes are 2-2.5 mm in length and have small tuft at base. Populus fremontii subs. Mar Aug Capsules contain seeds fremontii (Gulmon 1992, with long, silky Stettler, 1996, Clarke hairs. et al. 2007, Kleinman, n.d.) Potentilla anserina Fruits are oval, flat, subs. pacifica (Walker and reddish-brown and 2005, Stevens, n. d., about 2 mm in length. Baldwin et al. 2012) Pseudognaphalium Oblong fruits with californicum bristly, tuft-like (Keeley and Keeley projections (shed at 1987, Nesom 2013) maturity). Rosa californica Jul Sep Mature fruits (rose (Young 2001i, Lady hips) are bright red. Bird Johnson Each hip contains Wildflower Center multiple seeds. 2007) Salicornia bigelovii Sep Nov 1-1.5 mm curved, hairy (Glenn et al. 1997, seeds. Zedler 2001, Baldwin et al. 2012) Salicornia pacifica Oct Nov Mature seeds pinkish (Khan and Weber 1986, white, puberulent, and Young 200 lj) 0.5-1 mm long. Sequentially hermaphroditic. Salix exigua subs, May July Glabrous ovular exigua (Young and capsules. Small seeds Clements 2003, with pappus. Normally Anderson 2006, Clarke dispersed via wind or et al. 2007) water. Salix lasiolepis May Glabrous ovular (Bonner and Karrfalt capsules. 2008, Don 2014) Salvia apiana July Aug Shiny, light brown (Stevens 1994, fruit. Fruits are Montalvo and Beyers 2.5-3 mm in length. 2010d, Baldwin et al. 2012, Native Plant Database 2015) Salvia mellifera Jun Aug Dry calyces are gravity (Montalvo and Beyers dispersed. Up to 4 2010e) seeds/calyx. Seeds are 1 mm by 2 mm. Sclioenoplectus acutus Aug Sep Wide, smooth fruits with var. occidentalis 2 or 3 distinct sides. (Lacroix and Mosher Fruit 2-3 mm long and 1995, Johnson 2004, 1.2-1.7 mm wide. Baldwin et al. 2012, Baskin and Baskin 2014) Schoenoplectus 2-sided, smooth fruits. californicus Fruits 1.8-2.2 mm long (Stevens 2003, and 1.3 mm wide. Baldwin et al. 2012) Spartina foliosa Sep Nov 10-25 mm spikelets. (Zedler 2001, Baldwin et al. 2012) Stipa cernua Jul Aug Linear, smooth, glabrous (Laude et al. 1952, seed. Amme 2003, Herrera et al. 2006) Stipa lepida Brown fruit. Dark seeds (Ashby and Hellmers 4-7 mm in length. 1959, Elkhom Slough National Estuarine Research Reserve 2001, Amme 2003, Dave's Garden 2015b) Suaeda esteroa Oct Dec Two types of seeds: (Zedler 2001, Baldwin seeds can be et al. 2012) lenticular, black, and shiny (0.8-1.7 mm in length) or horizontal and matte (1-1.5 mm in length). Suaeda nigra Sep Oct Small, lenticular, shiny (Borders, n.d.) black seeds 0.5-2 mm long. Seed coat can be smooth, finely dotted, warty, net-like, or prickly. Suaeda taxifolia Jun Jul 1-2 mm horizontal or (Zedler 2001, Baldwin vertical seeds. Seeds et al. 2012) are shiny, lenticular and range from black to brown. Triglochiit maritima Jul Sep Mature inflorescences (Young 2002, Baldwin are brown. 1 seed/ et al. 2012, Recon fruit. Seeds of genus Native Plants Inc. usually linear. Seeds 2015) can be flat or angled. Vulpia microstachys 5.5-10 mm spikelets (Young and Young 1986, contain 4-6 mm long Howard 2006, Baldwin fruits. et al. 2012) Scientific name Seed collection details Abronia maritima Plants seed throughout the year, (Drennan 2008, Baldwin majority of seed production et al. 2012) occurs in late spring/summer. Removal of woody capsules aided with the use of generic nail clippers. Abronia umbellate Plants seed throughout the year, (Drennan 2008, Baldwin majority of seed production et al. 2012, Center occurs in late spring/summer. for Plant Conservation Removal of woody capsules 2015,) aided with the use of generic nail clippers. Achillea millefolium Cut entire inflorescences, collect (Baskin and Baskin in paper bags. Clean seeds with 2002a, Baldwin a hammermill, screen, and et al. 2012) fanning mill. Acmispon glaber Strip ripe seed pods from stems (Montalvo and Beyers by hand. Avoid breaking seeds 2010c) during thrashing. Rub pods with wooden block over #16 (medium) screen. Remove seeds from pods. Remove excess chaff with seed blower. Ambrosia psilostachya (Pavek 1992, Baldwin et al. 2012) Artemisia californica Strip brown inflorescence (Hauser 2006, Young- by hand. Mathews 2010, Baldwin et al. 2012) Artemisia douglasiana Seed is ready to harvest when it Elkhom Slough National can be easily removed from the Estuarine Research heads by shaking. Clip seed Reserve 2001, stalks and air dry in a paper Shultz 2014) bag. To thresh seeds rub the inflorescence through a screen. Remove chaff with a blower. Artemisia tridentate Seed from genus is ready to (Elkhom Slough harvest when it can be easily National Estuarine removed from the heads by Research Reserve 2001, shaking. Clip seed stalks and Baldwin et al. 2012, air dry in a paper bag. To thresh Tilley et al., n.d.) seeds mb the inflorescence through a screen. Remove chaff with a blower. Arthrocnemum Best to collect in November. subterminale (Zedler Collect inflorescences and air 2001, Clarke et al. dry. When dry, shake seeds 2007, Baldwin et al. from stalks. 2012) Astragalus pycnostachyus Other plants in genus, specifically var. ianosissimus A. sinuatus, have seeds that (McCue 2010, Baldwin mature in late July. et al. 2012, U.S. Fish and Wildlife Service, n.d.) Astragalus tetter var. Endangered species. Extract seeds titi (Showers 2010, from fruits by hand. Thresh seeds Baldwin et al. 2012) over sieve large enough to let set seed pass through. Run seeds through seed blower to remove parasitized or aborted seed. Atriplex calijarnica Gently seeds rub over # 18 sieve. (Young 2001a) Remove as much chaff as possible with a seed blower. Atriplex canescens Strip seeds from branches by (Springfield 1970, hand. If available, use a Baldwin et al. 2012) hammermill and a fanning mill to de-wing and clean seed. Collections made later (Dec-Apr) tend to have higher germination rates. Atriplex lentiformis Dioecious. (Young et al. 1980, Baldwin et al. 2012) Atriplex prostrated Fully mature fruit can be shaken (Khan and Ungar 1984, or hand stripped from Zedler 2001, Baldwin branches. Seeds will often et al. 2012) remain on bushes until April, so late collections are possible. Atriplex watsonii (Zedler 2001, Bryant 2004) Baccharis pilularis Dioecious. Collect seed heads by (Bonner and Karrfalt hand into open breathable bags. 2008, Montalvo et al. Alternatively shake branches over 2010b, Baldwin et al. a tarp. Fruit should be spread out 2012) to dry in a well-ventilated room or in the sun. Rub dried heads between palms or over a screen to remove the pappus and phyllaries. Baccharis salicifolia Dioecious. Collect ripe fruits by subs. Salicifolia hand or by shaking seeds onto (Bonner and Karrfalt canvases/tarps. Dry seeds at 2008, Baldwin et al. room temperature. Once dry, 2012) rub seeds over a screen to remove the pappus. Baccharis sarothroides Dioecious. Seeds can be collected (Bonner and Karrfalt by hand or branches can be 2008, Baldwin et al. shaken above tubs/tarps. 2012) Bads maritime Collect when fruits mature and (Zedler 2001, Marcone turn from green to white. 2003, Francis 2009, Extract seed from fruit. Dried Lonard et al. 2011) fruits should fragment easily, exposing seed Cressa truxtilensis Produces mature seeds from late (Elkhom Slough summer into early autumn. National Estuarine Research Reserve 2001, Zedler 2001) Croton californicus Dioecious. Collect July 15th- (Young 2001b, Baldwin November 17th. Remove chaff et al. 2012) by hand. Remove seeds from pods. Distichlis lift oralis Dioecious. Strip flowers with (Zedler 2001, Clarke seeds from inflorescences. et al. 2007, Baldwin et al. 2012) Distichlis spicata Dioecious. Seed is 2 mm long (Baskin and Baskin and brownish-gray at maturity. 2002b, Elsey-Quirk Rub seeds over #18 sieve to et al. 2009, Baldwin clean. et al. 2012) Encelia californica Achenes are wedge-shaped and (Bonner and Karrfalt densely compressed. Edges are 2008, Baldwin et al. long-ciliate and faces are 2012) glabrous or short-hairy. Collection timing is critical as achenes are easily blown from plant after reaching maturity. Eriogonum fasciculatum Best to collect from Jun-Jul. (Zedler2001, Montalvo Collect inflorescences as they and Beyers 2010a, begin to turn rusty brown. Push Baldwin et al. 2012) seeds through a screen to remove chaff. Fran ken ia salina Collect: September 16th- October (Young 2001k) 21st. Collect mature flowers and rub over #25 sieve. Use gloves when handling, the plant can be spiky. Grindelia camporum Harvest seed in June and again in (Zafar and Shah 1994, October. Clip seed heads or Bliss 2012) shake/rub mature seeds from seed heads into a collection bag. To clean, rub seed heads over sieve. Remove chaff using additional sieves or an air separator. Air dry in oven at 203 [degrees]F. Hazardia squarrosa (Keil et al. 2013) Heteromeles arbutifolia Clip or strip fruits from branches (Bonner and Karrfalt when bright red. Soak berries in 2008, Baldwin et al. water to ferment (over-soaking 2012, Gordon 2014, can be damaging). Pulp should Recon Native Plants float, making it easier to separate Inc. 2015) seeds from pulp. Alternatively, pulse berries in blender and then mb mixture over a screen to isolate fruit. Dry seeds before storing. RECON suggests keeping fruit intact. Hordeum brachyantherum Seed easily removed when stalks (Elkhom Slough are hand stripped. No National Estuarine additional cleaning required. Research Reserve 2001, Young 2001d) Isocoma menziesii Collect achenes golden in color, as (Zedler 2001, Wall and seeds are usually eaten by time Macdonald 2009, achenes turn brown. Shake ripe Montalvo and Beyers heads over open containers to 2010b) collect achenes. Alternatively, remove ripe heads and keep in porous bags. For 7. acradenia, Wall and Macdonald recommend rubbing flowers over a large screen, using a seed blower, and sieving over a #18 screen to separate seeds from bracts. Iva axillaris Strip seeds by hand or beat into a (Montalvo et al. hopper/open container. Rub 2010b) flower material over a screen and run through a blower to remove chaff. Jaumea carnosa Collect seed while fruits are (Young 2001e) swollen and green. Rub seeds over # 12 sieve to clean. Juncus acutus subs. leopoldii (Zedler2001, Baldwin et al. 2012) Juncus bufonius Seed capsules dehisce; seeds (Zedler 2001, Baldwin should be collected quickly et al. 2012) after plant death. Shake mature flowers to collect tiny seed. Limonium californicum Collections made in October are (Young 200If) best. Collect entire flower heads, which should detach easily when ripe. Rub flower heads over #20 sieve. Lupinus chamissonis Remove seeds from receptacles, (Young 200 lg) no further cleaning required. Lycium californicum Pick by hand. Berries best (Zedler 2001, Baldwin collected within 2 weeks of et al. 2012) appearance, otherwise birds will eat majority. Extract seeds from berries within a week, before berries begin to mold. Melica imperfecta Strip inflorescences. (Ashby and Hellmers 1959, Emery 1988, Young 2001h, Baskin and Baskin 2002c) Mimulus aurantiacus Rub seed capsules over a sieve. (Young 2001c, Baldwin et al. 2012) Oenothera elata Collect seed from spring cultivars (Greiner and Kohl in October and from winter 2014, B & T World cultivars in September. Bag Seeds 2015, Dave's seed heads and allow them to Garden 2015a, dry on plant or collect early and Kleinman, n.d.) allow to ripen in paper bags. Peritoma arborea Flowers several times/year (except (Lippitt et al. 1994, Dec-Jan). Ready for collection Borders et al. 2008, when capsules turn brown and Baldwin et al. 2012) are crisp. Strip mature fruits from plants by hand. Break apart pods by hand or with a hammermill or coater blender. Phacelia ramosissima Collect seed when flowers are dry (Baldwin et al. 2012) and brown. Strip seed from mature inflorescences directly into collection bag. Plantago erecta Dehiscent, ballistic seed dispersal. (Gulmon 1992, Montalvo Collect inflorescences into a et al. 2010a, Baldwin paper bag and let dry. Use sieve et al. 2012) to clean. Platanus racemosa Collect seedpods after they have (Bonner and Karrfalt turned brown. The task is 2008) easiest after leaves have fallen. Seedpods remain on trees into spring. Cut seedpods directly from tree. Crush dried seedpods to open. Remove dust and fine hairs. Populus fremontii subs. Dioecious. Collect seed as it is fremontii (Gulmon 1992, released from capsules during Stettler, 1996, Clarke dehiscence or collect entire et al. 2007, Kleinman, catkins prior to dehiscence. n.d.) Separate cotton fibers from seed. Potentilla anserina Let seeds dry on plant prior to subs. pacifica (Walker collection. 2005, Stevens, n. d., Baldwin et al. 2012) Pseudognaphalium californicum (Keeley and Keeley 1987, Nesom 2013) Rosa californica Collect hips as soon as they are (Young 2001i, Lady ripe. Extract seeds by hand from Bird Johnson dried hips. Alternatively, Wildflower Center macerate hips in water; remove 2007) floating seeds. Salicornia bigelovii Entire inflorescences should be (Glenn et al. 1997, collected and air-dried. When Zedler 2001, Baldwin dry, strip seeds from et al. 2012) inflorescences. Salicornia pacifica Collect inflorescences when plant (Khan and Weber 1986, tips are purple. Dry seeds on a Young 200 lj) screen for up to 3 months. Salix exigua subs, Dioecious. Harvest when catkins exigua (Young and are yellow-brown and capsules Clements 2003, begin to open. Shake catkins to Anderson 2006, Clarke remove dried seeds. et al. 2007) Salix lasiolepis Dioecious. Hand-harvest catkins (Bonner and Karrfalt when they begin to turn yellow/ 2008, Don 2014) brown. It is recommended to wait until capsules open. Separate seeds from cotton. Salvia apiana Collect seeds as capsules begin to (Stevens 1994, dry, before seeds are dispersed. Montalvo and Beyers Shake seeds from seed heads. 2010d, Baldwin et al. Use a sieve to isolate seeds. 2012, Native Plant Database 2015) Salvia mellifera Collect after inflorescences with (Montalvo and Beyers calyces are dry and brown. 2010e) Collect mature seeds by clipping, stripping, or shaking seed heads. Seed should be dried and passed through a sieve. Use of a blower is recommended. Sclioenoplectus acutus Because they are easily dispersed var. occidentalis by wind, it is important to (Lacroix and Mosher collect seeds close to the time 1995, Johnson 2004, of maturity. Seeds must be Baldwin et al. 2012, separated from the panicle and Baskin and Baskin cleaned. 2014) Schoenoplectus Harvest seed by hand from seed californicus heads. Alternatively, use shears (Stevens 2003, to clip entire seed heads from Baldwin et al. 2012) plant. Clean seeds. Spartina foliosa Multiple harvests may increase (Zedler 2001, Baldwin probability of collected good et al. 2012) seeds prior to dispersal or herbivory loss. Stipa cernua Harvest by hand or with a flow-vac (Laude et al. 1952, or combine at maturity. Collection Amme 2003, Herrera possible for 2-3 weeks. et al. 2006) Stipa lepida Seeds mature in spring. Allow (Ashby and Hellmers seed to mature on plant. At 1959, Elkhom Slough maturity, harvest seed. Clean National Estuarine prior to storage. Research Reserve 2001, Amme 2003, Dave's Garden 2015b) Suaeda esteroa Best to collect in Nov or early (Zedler 2001, Baldwin Dec. Cut whole inflorescence et al. 2012) or strip inflorescence. After cleaning, seeds should be dried. Suaeda nigra Collect when seeds are hard, (Borders, n.d.) black, and shiny when calyces will be brown and crumbly. Strip seeds from stalk by hand. Pass seed through a hammer mill or a sieve prior. Seeds should be spread out to dry before being processed/stored. Suaeda taxifolia Strip inflorescence by hand. (Zedler 2001, Baldwin et al. 2012) Triglochiit maritima Collect seeds between July 17-Sept. (Young 2002, Baldwin 23rd. Rub dry fruits between et al. 2012, Recon fingers to extract the seeds. Native Plants Inc. 2015) Vulpia microstachys Unknown exactly when seeds (Young and Young 1986, from S. California plants Howard 2006, Baldwin mature (intermountain varieties et al. 2012) mature late July-late September). Longevity Scientific name Seed germination (yrs.) Abronia maritima In the most successful 3 (Drennan 2008, Baldwin trials, achenes removed et al. 2012) from anthrocarp. Place achenes on filter paper in sterile petri dishes with ethephon or other 10-100 umol ethylene source. Incubate achenes in a chamber with alternating 12 h periods of light (27[degrees]C) and dark (20[degrees]C). Requires a sandy substrate. Abronia umbellate In the most successful 3 (Drennan 2008, Baldwin trials, achenes removed et al. 2012, Center from anthrocarp. Some seed for Plant Conservation lots require cold 2015,) pre-treatment. Germination requirements may differ year to year. For best results, sow clean seeds in the top 1" of a sandy growing medium. Achillea millefolium Lightly cover seeds with 3-5 (Baskin and Baskin growing medium (milled 2002a, Baldwin sphagnum, peat, perlite, et al. 2012) vermiculite w/ oscmocote). 90-100% germination rate. Acmispon glaber Heat or mechanical Long-lived (Montalvo and Beyers scarification needed to 2010c) break dormancy. Soak seeds in boiling water or heat in 120[degrees]C oven for 5 minutes for highest yield. Ambrosia psilostachya 3-5 (Pavek 1992, Baldwin et al. 2012) Artemisia californica Seeds will germinate when 2-5 (Hauser 2006, Young- fresh. Stored seeds need Mathews 2010, Baldwin to be exposed to light and et al. 2012) may require cold stratification. Other sources imply that pre-germination treatment is not necessary. Artemisia douglasiana Germinates naturally at 2-5 Elkhom Slough National relatively cool temps. Estuarine Research Reserve 2001, Shultz 2014) Artemisia tridentate No pre-germination treatment 2-5 (Elkhom Slough necessary. National Estuarine Research Reserve 2001, Baldwin et al. 2012, Tilley et al., n.d.) Arthrocnemum Seeds are highly germinable. 3-5 subterminale (Zedler Germination promoted by 2001, Clarke et al. low salinities. 2007, Baldwin et al. 2012) Astragalus pycnostachyus Flard seed coat may require Long-lived var. ianosissimus scarification. (McCue 2010, Baldwin et al. 2012, U.S. Fish and Wildlife Service, n.d.) Astragalus tetter var. If stored for an extended Long-lived titi (Showers 2010, period of time, hard seed Baldwin et al. 2012) coat may require scarification to initiate germination. 95% germination success rate on 0.5% agar plates with 11 hours light at 20[degrees]C and 13 hours dark at 12[degrees]C. Atriplex calijarnica Pre-planting: soak in water 10 (Young 2001a) for 24 hours, rinse. 86% germination rate after sowing in peat moss, perlite, nutrients, gypsum, and dolomitic lime. Germination occurs after 10 days. Atriplex canescens Germination is inhibited by 5-7 (Springfield 1970, lack of aeration, but Baldwin et al. 2012) improved with de-winging. Sow in medium with high substrate moisture at low temperatures, ideally at 18-24[degrees]C in California. Early collections may benefit from a 60 min. soak in sulfuric acid or a pre-chill at 5[degrees]C for 12 weeks. Atriplex lentiformis Maximum germination between 3-6 (Young et al. 1980, 10-25[degrees]C. Baldwin et al. 2012) Atriplex prostrated Readily propagated from 10 (Khan and Ungar 1984, seed. In the field, Zedler 2001, Baldwin germinates in late spring. et al. 2012) Atriplex watsonii Readily propagated from 10 (Zedler 2001, seed, germinates and Bryant 2004) establishes easily in field. Baccharis pilularis Seeds germinate without 1 (Bonner and Karrfalt pre-treatment. Cool 2008, Montalvo et al. temperatures yield highest 2010b, Baldwin et al. germination percentage. 2012) Baccharis salicifolia No pre-treatment necessary. 1 subs. Salicifolia Light necessary for (Bonner and Karrfalt germination. 2008, Baldwin et al. 2012) Baccharis sarothroides Germinates well in wet 1 (Bonner and Karrfalt soils. 2008, Baldwin et al. 2012) Bads maritime Difficult to grow from seed. 2+ (Zedler 2001, Marcone Marcone had success 2003, Francis 2009, exposing seeds to natural Lonard et al. 2011) light conditions and using a nutrient-enhanced potting medium. No known dormancy requirements. Cressa truxtilensis 2 (Elkhom Slough National Estuarine Research Reserve 2001, Zedler 2001) Croton californicus Pre-planting: soak seeds for 3 (Young 2001b, Baldwin 24 hours in water, cold et al. 2012) stratify for 30 days. Should germinate 30 days after sowing. Distichlis lift oralis 4-5 (Zedler 2001, Clarke et al. 2007, Baldwin et al. 2012) Distichlis spicata Seed germination highest 4-5 (Baskin and Baskin after wet stratification/ 2002b, Elsey-Quirk a fluctuating inundation et al. 2009, Baldwin regime and with low et al. 2012) salinity (Elsey-Quirk, 2009). Soak in water for 24 hours before sowing. Establishes well at restoration sites. Encelia californica To break dormancy, pre-soak 2-5 (Bonner and Karrfalt seeds in water. 2008, Baldwin et al. 2012) Eriogonum fasciculatum Seeds germinate well in 2-5 (Zedler2001, Montalvo flats. Light improves and Beyers 2010a, germination rate. Sow in Baldwin et al. 2012) fall-early winter. Fran ken ia salina Seeds need no pretreatment. 0-2 (Young 2001k) Germination naturally promoted by low salinity and high temperatures in spring. Grindelia camporum Soak in water under 3-5 (Zafar and Shah 1994, continuous light OR use Bliss 2012) two-stage cold stratification at 32[degrees]F and 59[degrees]F in the dark (Zafar 1994) to pre-treat. Hazardia squarrosa 0-1 (Keil et al. 2013) Heteromeles arbutifolia Fresh seeds germinate 2 (Bonner and Karrfalt readily. Chill stored 2008, Baldwin et al. seeds for 3 months at 2012, Gordon 2014, 3-5[degrees]C prior to Recon Native Plants sowing. Seeds germinate Inc. 2015) well 23[degrees]C. Hordeum brachyantherum No pre-treatment required. 4-5 (Elkhom Slough Sow seeds in May. Seeds National Estuarine should germinate 21 days Research Reserve 2001, after sowing. Germination Young 2001d) rate: 60%. Isocoma menziesii No pre-treatment required. 1+ (Zedler 2001, Wall and Seeds germinate well in Macdonald 2009, flats. Montalvo and Beyers 2010b) Iva axillaris Generally exhibits low Short- (Montalvo et al. germination rates. lived 2010b) Scarification is not effective. Cold stratification may be (studies needed). Jaumea carnosa Seeds germinate readily in 1 (Young 2001e) moist soil. Juncus acutus subs. Grows readily from seed in 2-5 leopoldii moderate salinities. (Zedler2001, Baldwin Clones can be dug entire et al. 2012) and transplanted. Juncus bufonius Seeds germinate readily in 2-5 (Zedler 2001, Baldwin low salinity soils. et al. 2012) Limonium californicum Sow in April. Propagates 1 (Young 200If) readily from seeds or plugs. Lupinus chamissonis Scarify using sandpaper for Long-lived (Young 200 lg) 5 minutes. Then, soak in hot water over night (repeat for seeds that do not imbibe). Sow in growing medium mid-October. Should germinate after 3 days. Lycium californicum Soak seeds in water for at 1 (Zedler 2001, Baldwin least 12 hours, then et al. 2012) transferred to moist soil. Reported germination rates are low, 5-10%. Melica imperfecta Plant has irregular 4-5 (Ashby and Hellmers germination patterns and 1959, Emery 1988, a low documented Young 2001h, Baskin germination rate, 30%. and Baskin 2002c) Literature inconsistent, certain sources suggest soaking seeds overnight in fresh water and cold stratifying for 2 weeks in peat. Conversely, Emery 1988 feels no pre-treatment necessary. Constant light (i.e. 24-hour photoperiod) exposure is documented to speed flowering. Mimulus aurantiacus Sow seeds in August. No 2-5 (Young 2001c, Baldwin pre-treatment needed, 50% et al. 2012) germination rate. Oenothera elata Surface sow (1 mm deep) to 3-5 (Greiner and Kohl ensure sufficient light. 2014, B & T World Long-day conditions (16 Seeds 2015, Dave's hours of light/ 8 hours of Garden 2015a, darkness) should be Kleinman, n.d.) simulated in the greenhouse. Should germinate after 15-30 days. Peritoma arborea Species does not require 2-5 (Lippitt et al. 1994, high soil moisture to Borders et al. 2008, germinate. Baldwin et al. 2012) Phacelia ramosissima 2 (Baldwin et al. 2012) Plantago erecta Non-dormant, no 3 (Gulmon 1992, Montalvo pre-treatment needed. With et al. 2010a, Baldwin ample water, will et al. 2012) germinate from Sep-Dec with varying temperatures. Platanus racemosa Cold moist stratification 2 (Bonner and Karrfalt needed to break dormancy. 2008) Populus fremontii subs. Germination is most 1 fremontii (Gulmon 1992, successful at Stettler, 1996, Clarke 20-30[degrees]C with et al. 2007, Kleinman, adequate moisture. Seed n.d.) should not be covered with soil. Potentilla anserina Non-dormant. Seeds should be Short- subs. pacifica (Walker planted in full sun in lived 2005, Stevens, n. d., lightly packed soil. Keep Baldwin et al. 2012) soil moist. Pseudognaphalium Germination stimulated by californicum presence of charred wood (Keeley and Keeley or aqueous extracts of 1987, Nesom 2013) charred wood. Rosa californica Soak seeds in water 2-4 (Young 2001i, Lady overnight prior to sowing. Bird Johnson Seeds germinate slowly, Wildflower Center cold stratification helps 2007) speed process. Salicornia bigelovii Irrigate with seawater. Root 1 (Glenn et al. 1997, zone salinity (top 15 cm Zedler 2001, Baldwin of soil) should be kept at et al. 2012) a salinity of 70-75 g for high yields. Salicornia pacifica Variety S. utahensis grows 2+ (Khan and Weber 1986, best at 5% NaCl treatment Young 200 lj) and under temperature regime of 15-5[degrees]C. Salix exigua subs, Seeds are non-dormant. 3-4 exigua (Young and Optimal germination Clements 2003, temperatures: Anderson 2006, Clarke 2-15[degrees]C. et al. 2007) Salix lasiolepis Sow near soil surface. 3 (Bonner and Karrfalt Exposure to light 2008, Don 2014) increases germination rate. Salvia apiana Scarification and possibly 2-4 (Stevens 1994, stratification needed to Montalvo and Beyers break seed dormancy. Sow 2010d, Baldwin et al. seed in early fall. Seeds 2012, Native Plant may respond to light, so Database 2015) plant in surface soil (1/8-1/4" deep). After planting, soak flats in water. Salvia mellifera Physiological dormancy. 1-2 (Montalvo and Beyers Exposure to light or 2010e) components of fire (charred wood, smoke, KN03) may stimulate germination. Sclioenoplectus acutus Physiological dormancy. Cold 2 var. occidentalis stratification breaks (Lacroix and Mosher dormancy. Germination 1995, Johnson 2004, rates are low for the Baldwin et al. 2012, species due to the thick Baskin and Baskin pericarp of the achene. 2014) Germination rates increase with overwintering in a pond or water source. Pre-treat seed with 0.05% solution of sodium hypochlorite 5 days prior to sowing. Seeds germinate to a higher percentage when grown in light. Schoenoplectus Plant seeds 1/4" under the californicus soil surface. Keep soil (Stevens 2003, surface moist and at a Baldwin et al. 2012) temperature of 100[degrees]F. Spartina foliosa Best success after cold 4 months (Zedler 2001, Baldwin storage in freshwater. et al. 2012) Stipa cernua Overheating can kill (Laude et al. 1952, seedlings. Amme 2003, Herrera et al. 2006) Stipa lepida No pretreatment needed. (Ashby and Hellmers Constant light (i.e. 1959, Elkhom Slough 24-hour photoperiod) National Estuarine exposure is documented to Research Reserve 2001, speed flowering. Amme 2003, Dave's Garden 2015b) Suaeda esteroa Seedlings establish well at 2+ (Zedler 2001, Baldwin restoration sites. et al. 2012) Suaeda nigra Pre-chill recommended. 3 (Borders, n.d.) Suaeda taxifolia Seeds germinate readily with 3 (Zedler 2001, Baldwin freshwater irrigation. et al. 2012) Triglochiit maritima No pre-treatment required. 3-5 (Young 2002, Baldwin et al. 2012, Recon Native Plants Inc. 2015) Vulpia microstachys Seeds germinate w/o 4-5 (Young and Young 1986, pretreatment. Heating and Howard 2006, Baldwin litter do not increase et al. 2012) germination. * In instances where published information is insufficient to fill-in portions of the table, cells are intentionally left blank. * Seed longevity data from Recon Native Plants Inc. used to supplement information found in the literature where necessary
Michelle L. Barton (1) *, Ivan D. Medel (2), Karina K. Johnston (2), and Christine R. Whitcraft (1)
(1) California State University Long Beach
(2) Santa Monica Bay Restoration Commission
* Corresponding author: email@example.com
Table 1. Suggested field, lab, and greenhouse equipment for seed collection, cleaning, and germination. Field equipment Lab/greenhouse equipment Collecting bins/paper bags Sieves of varying sizes (500 um-2 mm) Ziploc bags Paper envelopes Pens/pencils/markers Freezer Paper clips/binder clips Refrigerator Field data collection sheet Oven Clipboard Growing medium * Background documentation Sterile petri dishes * (recommended) Mesh screens/sieves (+) Flydrogen peroxide ([H.sub.2][O.sub.2]) * Tarp(s) (+) Nail clippers * Gloves (+) Mothballs * Gardening shears (+) Ethylene (ethephon or sliced apple) * Jepson manual (+) * = species specific + = optional Table 2. General seed collection method based on plant anatomy (Wall 2009). Fruit/seed type Collection techniques Moist fruits/berries Hand-pluck fruits. Dehiscent species Collect entire inflorescences prior to dispersal. Alternatively, secure cloth bags around ripening stalks to capture dispersed seed. Inflorescences Strip inflorescences. Seed heads Shake ripe seed directly onto a tarp or collection bag underneath the target plant. Seed clusters Remove entire seed cluster from plant. Table 3. Detailed methodology for techniques commonly employed to break seed dormancy. Method General description Scarification Mechanically scar seed coat with sandpaper, knives, files, or clippers. Alternatively soak seed in acid or hot water (Emery 1988, Lippitt 1994, Bonner and Karrfalt 2008). Hot water treatment Place seeds into hot water (180-200[degrees]F) and leave them to soak as the water cools (Emery 1988, Bonner and Karrfalt 2008). Dry heat Expose seeds to 180-212[degrees]F heat. Use of an incubator, rather than oven, preferred (Emery 1988). Charate Expose seeds to ash from burned plants. This may neutralize germination inhibitors in species that naturally germinate when exposed to fire (Emery 1988, Baskin and Baskin 2014). Fire Expose seeds to direct flame. This may be effective as a means to spur germination in species that naturally germinate when exposed to fire (Baskin and Baskin 2014). Water Soak seeds in water to leach out water-soluble inhibitors (Baskin and Baskin 2014). Cold stratification Store seeds in cold conditions (35-41[degrees]F) for 1 -3 months to simulate winter conditions (Bonner and Karrfalt 2008, Elsey-Quirk et al. 2009, Baskin and Baskin 2014). Warm stratification Store seeds in warm conditions (65[degrees]F or higher) (Baskin and Baskin 2014).
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|Author:||Barton, Michelle L.; Medel, Ivan D.; Johnston, Karina K.; Whitcraft, Christine R.|
|Publication:||Bulletin (Southern California Academy of Sciences)|
|Date:||Apr 1, 2016|
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